|Scientific Name:||Lama guanicoe|
|Species Authority:||(P.L.S. Müller, 1776)|
|Taxonomic Notes:||Although four Guanaco subspecies have been historically described, based on morphological traits such as skull and body size and hair color (Wheeler 1995; González et al., 2006), no further studies based on large sample sizes have been conducted to account for morphological variation. These subspecies were associated to four geographic areas divided mainly by the Andean Cordillera (Franklin, 1982; Torres, 1992). According to this classification, L. g. cacsilensis (Lönnberg, 1913) would occupy Peru, northern Chile and part of the highlands of Bolivia between 8° and 22° S; L. g. voglii (Krumbiegel, 1944) would spread across the northwest of the Bolivian Chaco region, Paraguay and the Chacoan Argentina from 35° S northwards; L. g. huanacus (Molina, 1782) across northern and central Chile between 22° and 38° S; and L. g. guanicoe (Müller, 1776) throughout western Argentina spreading southward across the Argentine and Chilean Patagonia and Tierra del Fuego and Navarino islands, and eastward until the Atlantic coast.
More recently, molecular evidence has shown that there is no variation among the proposed subspecies and that Guanacos are a monophyletic group and the ancestors of the domestic Llama; whereas Vicuñas are ancestors of the domestic Alpaca (Kadwell et al. 2001). However, the northern populations of Peru and northern Chile showed some degree of differentiation from the populations in the Bolivian Chaco, Argentina, and central and southern Chile (Marín et al. 2008), suggesting that only two subspecies would exist: L. guanicoe cacsilensis, found in the north from 8° to 20°S, and L. guanicoe guanicoe, found in the south from 22° to 55°S (Marín et al. 2006, Wheeler 2006).
|Red List Category & Criteria:||Least Concern ver 3.1|
|Assessor(s):||Baldi, B., Lichtenstein G., González, B., Funes, M., Cuéllar, E., Villalba, L., Hoces, D. & Puig, S.|
|Reviewer(s):||Baldi, R. & Lichtenstein, G. (South American Camelid Red List Authority)|
This species is considered to be Least Concern due to a wide distribution, presumed large populations and occurrence in numerous protected areas. Guanacos remain a conservation dependant species despite, and although they do not meet any threat criteria they remain a research and management priority. It is important to stress that the future of this species depends on the implementation of conservation-oriented management at local, national and international levels. Poaching continues to be intense. Also, the increasing demand to develop shearing initiatives could impose a serious risk to local and even regional populations if the effects of this type of management are not properly accounted for. As the demand to use guanacos is concentrated on the few scattered high-density populations, most guanacos might be affected by intensive use in the near future. Today, around 35% of the high density populations identified in Argentina are under the effect of shearing initiatives Failure in assessing the effects of management and in developing ecologically sustainable management actions can result in a rapid population decline, especially in southern Argentina where the species is more abundant. At the national level, guanacos are likely to become extinct in three out of the five countries comprising their historic distributional range.
|Range Description:||Guanaco is a widespread species with an extensive, though discontinuous, range from the north of Peru to Navarino Island in southern Chile. Most of guanacos are found in Argentina. Although its range covers most of Patagonia, guanaco populations appear to be more scattered towards the north of the region (Chubut, Río Negro, Neuquén and southern Mendoza provinces) than in Santa Cruz and Tierra del Fuego (Walker et al., unpublished data). Through northern Patagonia, guanaco distribution extends highly fragmented in relict populations in La Pampa and southwestern Buenos Aires provinces. Through central and northern Argentina guanaco distribution is restricted to the western half of the country, along the pre-Andes and Andean mountains up to the border with Bolivia (Baigún et al. 2007), although a relict population has been recently reported in the arid Chaco of northwestern Córdoba (Schneider et al. unpublished data). In Chile, the largest guanaco populations are concentrated in the Magallanes and Aysén regions in the south (with the highest numbers on the Tierra del Fuego island) and Tarapaca region in the north. In Bolivia, a relict population of guanaco persists in the Chaco region (Cuéllar and Fuentes 2000) and recent observations of guanacos have been reported in the southern highlands between Potosi and Chuquisaca (Nuñez, unpublished data). Although Pinaya (1990) reported the presence of guanacos in southeastern Tarija, these records need to be confirmed. In Paraguay, a relict population has been reported in the northwestern Chaco (Villalba 2004). Peru is the northernmost part of guanaco distribution at approximately 8°30'S, at the Calipuy National Reserve in La Libertad department. To the south, populations reach the Salinas Aguada Blanca National Reserve in Arequipa and Moquegua departments (16°10'S). A guanaco population was recorded in the Nevado Salcantay area, in the Anta district (Wheeler 2006, Veliz and Hoces 2007).|
Native:Argentina; Bolivia, Plurinational States of; Chile; Paraguay; Peru
|Range Map:||Click here to open the map viewer and explore range.|
Although in the Argentine Patagonia guanaco distribution is rather continuous, densities are typically low (<5 guanacos/km²) or very low as in many parts of Chubut, Rio Negro and Neuquén (<2 guanacos/km²) while higher density populations are scarce and spatially structured (Baldi et al. 2001, Baldi et al. in press, Novaro et al. 2007, Puig et al. 1997, 2003). A global estimate for continental Patagonia was reported by Amaya et al. (2001) after aerial censuses, but the numbers are used as a reference as the errors associated to sampling were too large to be conlclusive about the results. For the rest of Argentina, population densities are below 1 guanaco/km² and the population is highly fragmented (Baigún et al. 2007, Puig and Videla 2007), with some relict populations in La Pampa, Córdoba and Buenos Aires provinces. Other high-density populations occur in southern Chile reaching up to 43 guanacos/km² at Torres del Paine National Park (Sarno and Franklin 1999) but in the rest of the range populations are small and highly scattered.
The global estimate for the guanaco population is below 600,000 animals, and almost 90% of the population is found in Argentina. However, differences in survey methodology and effort invested across such a vast area make necessary to be cautious about the figures, and these should be taken as a reference. In particular, a more reliable estimate is needed for the continental Argentine Patagonia, as most of the guanacos are found in that region and its number greatly affects global population size. Also for Chile the estimate is rather speculative as it results from scattered information instead of planned surveys (B. González, unpublished data). As a general rule, it is recommended to use the distance sampling method, either for ground or aerial surveys, as it is based on more realistic assumptions than the fixed-width strip transect methods which tend to underestimate population numbers (Buckland et al. 2001). However, where numbers are too low as in relict populations, total counts or less systematics methods can be appropriate. Also, extrapolation of local densities to larger areas must be careful and made according to sampling effort. Accurate estimates of local densities are not sufficient at the time to estimate abundance for larger areas unless the sampling effort is properly disseminated throughout the region.
Country: Guanaco population
|Habitat and Ecology:||
Most of the guanacos are distributed across the phytogeographic provinces of the Monte and Patagonia, arid and semi-arid shrublands and grasslands comprising over 700,000 km². Guanacos are also found in the Puna, pre-Puna, Andean steppe, Chacoan grasslands and shrublands, Espinal and the southern Pampa. There are both migratory and sedentary populations across the guanaco range.
Adult guanacos weigh 80-120 kg and their breeding system is a resource defense polygyny. An adult male defends a territory where a group of females and their offspring (chulengos) feed, from the intrusion of other males (Franklin 1982, 1983). In a recent study, it was shown that vigilance and foraging accounted for almost 90% of the diurnal time budget of territorial male and female guanacos, but guanacos benefited from living in groups as individual foraging time increased with group size as well as collective vigilance against pumas, suggesting that predation risk is a strong component modeling guanaco social structure (Marino and Baldi 2008).
Guanacos are generalist herbivores of intermediate selectivity (i.e. they include a large proportion of both grasses and shrubs in their diets) (Raedeke and Simmoneti 1988, Fraser 1998, Puig et al. 1997, 2001, Baldi et al. 2004). The domestic sheep was the main ungulate species introduced across the guanaco range, reaching 22 million heads within 50 years in the Argentine Patagonia. Both guanacos and sheep largely overlap in their forage preferences. Over 80% of their diets are identical (Puig et al. 2001). Although both species can include over 100 plant species in their diets, only 17 species made up 80% of the diets, and in Patagonia two grass species represented 40% of both guanaco and sheep diets (Baldi et al. 2004).
Across the extensive Patagonian rangelands, competition with sheep, hunting, and habitat modification have resulted in guanacos occupying marginal, low quality lands in terms of vegetation cover and the availability of the most important plant species in their diet, since sheep ranching monopolized the most productive areas (Baldi et al. 2001). A preliminary study in the Bolivian Chaco showed that the guanaco is a generalist feeder, responding to the seasonal availability of fruits, flowers and leaves, including a variety of cacti (Cuéllar, unpublished data). In this region guanacos compete for forage and spatial resources mainly with cattle and horses (Cuéllar, unpublished data).
|Use and Trade:||
80% of the total population is utilized (20% legally and the remainder illegally).
A proportion of fibre from guanaco farms results from individuals captured in the wild as newborns and raised in captivity, while other proportion of the fibre comes from capturing, shearing and release of individuals in wild populations (wild management).
Guanacos are still numerous and widely distributed but continued the precipitous decline initiated in the 19th century. Over-hunting, range degradation either due to overstocking with domestic livestock or extractive activities, and interspecific competition for forage have all played a significant role in the demise of guanacos all across the distributional range (Raedeke 1979; Franklin 1982, Miller et al. 1983, Cunazza et al. 1995; Cuellar and Fuentes 2000, Puig et al. 2001, Baldi et al. 2001; Baldi et al. 2004). In Bolivia the current major threat is habitat loss after the sport hunting was halted in 2001 (Cuéllar, unpublished data). In Chile, recreational hunting and poaching are major threats, while in Argentina they are still widely spread. Also in Peru guanacos are seriously affected by poaching and subsistence hunting. In addition, health studies conducted in mainland Patagonia have shown that guanaco populations are relatively disease-free but are themselves susceptible to common diseases of domestic livestock (sheep, cattle, and horses) (Karesh et al. 1998; Beldomenico et al. 2003; Uhart et al. unpublished. data). Castillo (2006) came to a similar conclusion based on the study of parasite load in free ranging Peruvian guanacos.
Today the guanaco occupies only 40% of its original range (Puig 1995; Franklin et al. 1997) and the distribution has become fragmented into smaller, relatively isolated populations. Although the species is not threatened with demographic extinction at a continental scale, it is ecologically extinct in most of its remaining range (Novaro et al. 2000), and some southern populations are under serious risk of local or even regional extirpation (Cunazza et al. 1995), and it has been predicted that the northern subspecies L. guanicoe cacsilensis will become extinct in Peru within 30 years if current hunting off-take rates are not curtailed (www.conopa.org). Spatial fragmentation is a threat to guanaco populations. Human activities such as hunting, mining, oil exploration and extraction, fencing, and the development of infrastructure and habitat loss often impose barriers to animal movements and prevent travel by individuals between populations. The loss of connectivity results in small, closed and isolated populations under increasing risk of collapse due to either loss of genetic variation, environmental or demographic stochasticity – the latter highly relevant to the very small populations in the Chaco region (Cuéllar et al. 2001). Recent findings are showing that low genetic variation may lead to reproductive failure and congenital malformations (Franklin and Grigione 2005, Zapata et al. 2008, Marin et al. in preparation).
Increasing pressure by private landowners in Patagonian rangelands may result in a threat to the remaining high-density guanaco populations if management is not properly planned and implemented. Live-shearing and subsequent release of wild guanacos could contribute to their conservation only if the effects of this activity are properly assessed and management is applied accordingly. If not ecologically sustainable, the viability of the most important guanaco populations will be taken into risk and hence the global population might collapse. Since 2003, around 11,000 guanacos have been captured at 7 ranches in just one Patagonian province (Baldi et al. in press). Careful evaluation of current management practices involving live shearing is urgently needed.
Land desertification due to overgrazing coupled with more severe and frequent droughts associated to climate change are threats requiring urgent attention as they can have global, major effects on guanaco abundance throughout the range. Severe droughts can have drastic effects on local guanaco populations as it was documented for eastern Patagonia (R. Baldi, unpublished data). In addition, models on climate change predict a sharp decrease in rain precipitation within the next 50 years in the arid Southern South America (Nohara et al. 2006). Therefore, it is crucial to favour the ecological functionality of guanaco populations through adequate management as a step to mitigate additional effects of climate change in the near future.
The species is found in a number of protected areas and is included in the appendix II of CITES, thus regulating international trade of products derived from the guanaco.
Chile: Species protected by the Ley de Caza, Criaderos y Uso in situ(Law on hunting, breeding and in situ use). However, personnel for law enforcement are insufficient. Only 4% of the habitats of guanaco have effective protection (8354 sq. km., in 8 National Parks and 4 Reserves). Additionally there are fiscal and private areas in which hunting is prohibited, either with relict populations (7750 sq. km.), or to protect the species (1212 sq. km.). A National Management Plan does not exist.
Peru: Recent legislation ratifies the classification of the guanaco in Peru as "Endangered" as of 2004. Active management of populations is carried out by CONACS (Consejo Nacional de Camélidos Sudamericanos National Council for South American Camelids), and local communities.
Bolivia: A team of the Wildlife Conservation Society (WCS) is working with the government, local authorities and local people to strengthen the management of protected areas. The government has issued an official notice to law enforcement offices in the region concerning the protection of guanaco. The main conservation aim has been achieved thanks to the permanent presence of trained indigenous parabiologists in the remaining range of the species (Cuéllar, unpublished data).
Argentina: A National Management Plan (Plan Nacional de Manejo del Guanaco) has been prepared and endorsed by the provinces with the highest guanaco densities. This plan was coordinated by the Dirección de Fauna Silvestre (Federal Wildlife Agency), with the participation of various local institutions, and has a main focus towards Patagonia. The federal wildlife conservation law (Ley Nacional de Conservación de la Fauna) and various provincial acts provide a legal basis for the protection and use of the species. In Patagonia, guanaco conservation activities include sustainable use of the species in the wild, regulation of hunting quotas, and closing of some access routes and oil trails. Protected areas in the Patagonian steppe would encompass 10% of the guanacos if effective, but most of the protected areas are rather nominal as they contain livestock, there are no wardens and poaching is common. The percentage of the area under effective protection in the Patagonian steppe was estimated to be less than 1% (Walker et al. 2004). In the central provinces there are 11 protected areas (either national, provincial or private). However, the progress made in legislation and management tools, implementation of actions is needed.
Amaya, J. N., von Thüngen, J. and Delamo, D. A. 2001. Relevamiento y distribución de guanacos en la patagonia. Informe final. Comunicación Técnica Nº 111. Área Recursos Naturales, EEA-Bariloche/ GTZ. INTA EEA Bariloche – C.C. Nº 277 (8400), Bariloche, Río Negro, Argentina.
Baigún, R., Bolkovic, M. L., Aued, M. B., Li Puma, M. C., Scandalo, R., Nugent, P. and Ramadori D. 2007. Censo de Camélidos Silvestres al Norte del Río Colorado. Dirección de Fauna Silvestre. Secretaría de Ambiente y Desarrollo Sustentable, Buenos Aires, Argentina.
Baldi, R., Albon, S. D. and Elston, D. A. 2001. Guanacos and sheep: evidence for continuing competition in arid Patagonia. Oecologia 129: 561-570.
Baldi, R., Novaro, A., Funes, M., Walker, S., Ferrando, P., Failla, M. and Carmanchahi, P. In press. Guanaco management in Patagonian rangelands. In: Johan du Toit, Richard Kock and James Deutsch (eds), Conservation Science and Practice: Conserving Rangelands, Blackwell Publishing.
Baldi, R., Pelliza-Sbriller, A., Elston, D. and Albon, S. D. 2004. High potential for competition between guanacos and sheep in Patagonia. Journal of Wildlife Management 68(4): 924-938.
Beldoménico, P. M., Uhart, M., Bono, M. F., Marull, C, Baldi, R. and Peralta, J. L. 2003. Internal parasites of free-ranging guanacos from Patagonia. Veterinary Parasitology 118: 71-77.
Buckland, S. T., Anderson, D. R. Burnham, K. P. and Laake, J. L. 1993. Distance Sampling: Estimating Abundance of Biological Populations. Chapman and Hall, London, UK.
Castillo, V. H. 2006. Contribución al estudio del parasitismo gastrointestinal en guanacos (Lama guanicoe cacsilensis) silvestres del Perú. Resúmenes y Trabajos del IV Congreso Mundial sobre Camélidos.
Cuéllar, E. and Fuentes, A. 2000. Censo aéreo de guanacos Lama guanicoe en el Chaco cruceño. Revista Boliviana de Ecología y Conservación Ambiental 8: 83-90.
Cuéllar, E., Noss, A. and Cuéllar, R. L. 2001. Imminent Infamy? Wildlife Conservation Magazine, USA.
Cunazza, C., Puig, S. and Villalba, L. 1995. Situación actual del guanaco y su ambiente. In: S. Puig (ed.), Técnicas para el Manejo del Guanaco. IUCN, Gland, Switzerland.
Franklin, W. L. 1982. Biology, ecology, and relationship to man of the South American camelids. In: M. Mares and H. Genoways (eds), Mammalian biology in Soth America. Special publication series Pymatuning laboratory of ecology, pp. 457-489. University of Pittsburgh.
Franklin, W. L. 1983. Contrasting socioecologies of South America’s wild camelids: The vicuña and the guanaco. In: J. F. Eisemberg and D. G Kleiman (eds), Advances in the Study of Mammalian Behavior, pp. 573-629. The American Society of Mammalogists.
Franklin, W. L. and Grigione, M. M. 2005. The enigma of guanacos in the Falkland Islands: the legacy of John Hamilton. Journal of Biogeography 32: 661-675.
Franklin, W. L, Bas F., Bonacic, C. F., Cunazza, C. and Soto, N. 1997. Striving to manage Patagonia guanacos for sustained use in the grazing agroecosystems of southern Chile. Wildlife Society Bulletin 25(1): 65-73.
Fraser, M. D. 1998. Diet composition of guanacos (Lama guanicoe) and sheep (Ovis aries) grazing in grasslands communities typical of UK uplands. Small Ruminant Research 29: 201-212.
González, B. A., Palma, R. E., Zapata, B., Marín, J. C. 2006. Taxonomic and biogeographical status of guanaco Lama guanicoe (Artiodactyla, Camelidae). Mammal Review 36(2): 157-178.
IUCN. 2008. 2008 IUCN Red List of Threatened Species. Available at: http://www.iucnredlist.org. (Accessed: 5 October 2008).
Kadwell, M., Fernandez, M., Stanley, H., Baldi, R., Wheeler, J. C., Rosadio, R. and Bruford, M. 2001. Genetic analysis reveals the wild ancestors of the llama and alpaca. Proceedings of the Royal Society of London B Biological Sciences 268: 2575-2584.
Karesh, W., Uhart, M., Dierenfeld, E., Braselton, W., Torres, A., House, C., Puche, H. and Cook, R. 1998. Health evaluation of free-ranging guanaco (Lama guanicoe). Journal of Zoo and Wildlife Medicine 29: 134–141.
Krumbiegel, I. 1944. Die neuweltlichen Tylopoden. Zoologischer Anzeiger 145: 45–70.
Lönnberg, E. 1913. Notes on guanacos.
Marín, J. C., Spotorno, A. and Wheeler, J. 2006. Sistemática molecular y filogeografía de camélidos Sudamericanos: implicancias para su conservacion y manejo. In: B. Vilá (ed.), Investigacion, conservacion y manejo de vicuñas, pp. 85–100. Proyecto MACS Manejo Sostenible de Camélidos, Buenos Aires, Argentina.
Marín J. C., Spotorno, A. E., González, B. A., Bonacic, C., Wheeler, J. C., Casey, C. S., Bruford, M. W., Palma, R. E. and Poulin, E. 2008. Mitochondrial DNA variation and systematics of the guanaco (Lama guanicoe, ARTIODACTYLA: CAMELIDAE). Journal of Mammalogy 89(2): 172-184.
Marino, A. and Baldi, R. 2008. Vigilance patterns of territorial guanacos (Lama guanicoe): the role of reproductive interests and predation risk.
Miller, S. D., Rottmann, J., Raedeke, K. J. and Taber, R. D. 1983. Endangered mammals of Chile: status and conservation. Biological Conservation 25: 335-352.
Molina, G. I. 1782. Saggio Sulla Storia Naturale del Chili. Tomasso d’Aquino, Bologna, Italy.
Müller, P. L. S. 1776. Erste Classe, Säugende Thiere. Des Ritters Carl von Linné vollständiges Naturalsystem nach der zwölften Lateinischen Ausgabe, 1773–1776 36: 1–62.
Nohara, D., Kitoh, A., Osaka, M. and Oki, T. 2006. Impact of climate change on river discharge projected by multimodel ensemble. Journal of Hydrometeorology 7: 1076-1089.
Novaro, A. J., Funes, M. C. and Walker, R. S. 2000. Ecological extinction of native prey of a carnivore assemblage in Argentine Patagonia. Biological Conservation 92: 25-33.
Novaro, A., Walker, S., Funes, M., Radovani, N., Bolgeri, M., Puig, S. and Videla, F. 2007. Estrategia de conservación de guanacos en Patagonia norte, basada en la conectividad del paisaje. III. Reunión Binacional de Ecologia, La Serena, Chile.
Pinaya, I. 1990. Informe inspección ocular proyecto de Decreto Supremo, creación Reserva Biológica Cordillera de Sama. DVSPN y C. CDF, M.A.C.A. Marzo de 1990.
Puig, S. 1995. Abundancia y distribución de las poblaciones de guanacos. In: S. Puig (ed.), Técnicas para el Manejo del Guanaco, pp. 57-70. IUCN, Gland, Switzerland.
Puig, S. and Videla, F. 2007. Distribución, densidades y estado de conservación de los camélidos. In: E. Martínez Carretero (ed.), Diversidad Biológica y Cultural de los Altos Andes Centrales de Argentina. Línea de Base de la Reserva de Biósfera San Guillermo, pp. 199-223. Univ Nac San Juan, San Juan.
Puig, S., Ferraris, G., Superina, M. and Videla, F. 2003. Distribución de densidades de guanacos (Lama guanicoe) en el norte de la Reserva La Payunia y su área de influencia (Mendoza, Argentina). Multequina (Latin American Journal of Natural Resources) 12: 37-48.
Puig S., Videla, F. and Cona, M. 1997. Diet and abundance of the guanaco (Lama guanicoe Müller 1776) in four habitats of Northern Patagonia, Argentina. Journal of Arid Environments 36: 343-357.
Puig, S., Videla, F., Cona, M. I. and Monge, A. 2001. Use of food availability by guanacos (Lama guanicoe) and livestock in northern Patagonia (Mendoza, Argentina). Journal of Arid Environments 47: 291-308.
Raedeke, K. and Simonetti, J. 1988. Food habits of Lama guanicoe in Atacama desert of northern Chile. Journal of Mammalogy 69: 198–201.
Raedeke, K. J. 1979. Population dynamics and socioecology of the guanaco (Lama guanicoe) of Magallanes, Chile. Ph.D. Thesis, University of Washington.
Sarno, R. and Franklin, W. L. 1999. Population density and annual variation in birth mass of guanacos in southern Chile. Journal of Mammalogy 80: 1158-1162.
South American Camelid Specialist Group/Grupo Especialista en Camélidos Sudamericanos. Available at: http://www.iucn.org/themes/ssc/sgs/gecs.
Torres, H. 1992. South American Camelids: An Action Plan for their Conservation. IUCN/SSC South American Camelids Specialist Group, Gland, Switzerland.
Véliz, C. and Hoces, D. 2007. Distribución potencial del guanaco y de la vicuña en el Perú. XII Reunión del Seminario Permanente de investigación Agraria SEPIA. Tarapoto, Perú.
Villalba, L. 2004. Monitoreo de la población de Lama guanicoe voglii. Technical Report. DesdelChaco Foundation, Loma Plata, Paraguay.
Villalba, L. 2008. Informe Proyecto Guanacos. DesdelChaco Foundation, Loma Plata, Paraguay.
Walker, S., Novaro, A., Funes, M., Baldi, R., Chehebar, C., Ramilo, E., Ayesa, J., Bran, D., Vila, A. and Bonino, N. 2004. Rewilding Patagonia. Wild Earth 15: 32-37.
Wheeler, J. C. 1995. Evolution and Present Situation of the South-American Camelidae. Biological Journal of the Linnean Society, 52: 271–295.
Wheeler, J. C. 2006. Guanaco 1: Working to Save Peru’s Endangered Guanacos. Camelid Quarterly 5(1): 39-42.
Zapata, B., González, B. A., Marin, J. C., Cabello, J. L., Johnson, W. and Skewes, O. 2008. Finding of polydactyly in a free-ranging guanaco (Lama guanicoe). Small Ruminant Research 76: 220-222.
|Citation:||Baldi, B., Lichtenstein G., González, B., Funes, M., Cuéllar, E., Villalba, L., Hoces, D. & Puig, S. 2008. Lama guanicoe. The IUCN Red List of Threatened Species. Version 2014.3. <www.iucnredlist.org>. Downloaded on 01 April 2015.|